Onychophora have undergone a new wave of taxonomic revisions in the last three years. After a recent revision, only two species of Macroperipatus are registered from Brazil: Macroperipatus ohausi and Macroperipatus geayi. The former was only the second species of Onychophora to be described from Brazil, and is herein redescribed, illustrated, and its taxonomic status revised. Macroperipatus ohausi had only been collected at the type locality, Petrópolis, state of Rio de Janeiro, but has not been found there since its discovery. In 2009 we collected two specimens of this species in an Atlantic rain forest fragment in Nova Iguaçu in RPPN dos Petroleiros (also state of Rio de Janeiro). The color of the new specimens was interpreted and described based on the NBS/ISCC System of Color Designation, and corresponds only weakly to that described in earlier publications on M. ohausi. The identification of M. ohausi was based solely on its morphology and compared with its original description. In addition, a female syntype of M. ohausi deposited in the Zoologisches Museum (Hamburg) was examined. We reinvestigated its external morphology and concluded that the dorsal papillae have been misinterpreted in M. ohausi in previous studies of its integument. The new specimens examined are conspecific with M. ohausi mainly by having conical primary papillae, different sized accessory papillae, and by the number of legs. Our material resembles the original description of M. ohausi, for example, in spinous pad and nephridial tubercles, even though these features are not represented in the original description and rede- scription of M. ohausi; these features are further detailed here. We concluded that the dorsal papillae morphology of M. ohausi, shares diagnostic characters, for example, with representatives of the genera Peripatus and Epiperipatus. The dorsal papillae are rounded and not square as in the original description and redescription. Another argument that supports our conclusion is that the RPPN dos Petroleiros, the locality where the new samples were collected, belongs to the same mountain chain and Atlantic mosaic as the original material of M. ohausi. Furthermore, a male of M. ohausi is described here for the first time. Based on our morphological data, we transfered this species to the genus Epiperipatus, proposing a new combination, E. ohausi.
Onychophora experimentó una nueva serie de revisiones taxonómicas en los últimos tres años. Después de una reciente revisión, ]]>
Macroperipatus fueron registradas para Brasil: Macroperipatus ohausi y Macroperipatus geayi. La primera fue ]]>
Macroperipatus ohausi sólo se había recolectado en la localidad tipo, Petropolis, Estado de Rio de Janeiro, pero no se ha encontrado allí desde su descubrimiento. En 2009, recolectamos dos ejemplares de esta especie en un fragmento de la ‘Mata Atlantica’ en Nova Iguaçu en el RPPN de los Petroleiros ]]>
M. ohausi. La identificación de M. ohausi
se basó únicamente en su morfología, en comparación con la descripción original. Además, se examinó un sintipo hembra de M. ohausi depositado en Zoologisches Museum (Hamburgo). Nosotros reinvestigamos su morfología externa y concluimos que las papilas dorsales fueron mal interpretadas en
M. ohausi en los estudios previos de su integumento. Los nuevos especímenes examinados son coespecíficos con M. ohausi principalmente por tener las papilas primarias cónicas, diferente tamaño de las papilas accesorias, y por el número de patas. Nuestro material se asemeja a la ]]>
M. ohausi, por ejemplo, la almohadilla espinosa y tubérculos nefridiales, aunque estas características no están representadas en la descripción original y redescripcion de M. ohausi, estas ]]>
M. ohausi, comparte caracteres de diagnóstico, por ejemplo, con los representantes de los géneros Peripatus y Epiperipatus. Las papilas dorsales son redondeadas y no cuadradas como en la descripción original y redescripción. Otro argumento que apoya nuestra conclusión es que la RPPN dos Petroleiros, la ubicación en la que se recogieron las nuevas muestras, pertenece a la misma cadena de montañas y al mosaico Atlántico, tal como el material original de M. ohausi. Por otra ]]>
M. ohausi. Basado en nuestros datos morfológicos, transferimos esta especie al género Epiperipatus, que propone una nueva combinación, E. ]]>
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Palabras clave: gusanos de terciopelo, Peripatidae, taxonomía, distribución geográfica, Brasil.
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Onychophorans, commonly referred to as velvet worms, are a small group of terrestrial invertebrates that are predominantly found in soil, leaf litter, or rotten wood. The species of Onychophora are arranged in two large families, which display a Pangean distribution (Morera-Brenes 2012, Murrienne et al. 2014). Currently 16 species are known from Brazil, belonging to four genera within the family Peripatidae (
Peripatus Guilding, 1826, Epiperipatus Clark, 1913, Macroperipatus Clark, 1913, and Oroperipatus Cockerell, 1908). However, Sampaio-Costa, Chagas-Junior & Baptista (2009) recorded 24 morphospecies, of which half are undescribed. ]]>
The second oldest species described from Brazil is Peripatus ohausi Bouvier, 1900 col- lected in the city of Petrópolis (state of Rio de Janeiro). Bouvier (1900) gave a modest description of P. ohausi, but singular enough to distinguish it from other species known during that time. Its description was extended in Bouvier (1905) based only on the syntypes. Clark (1913) proposed the subgenus Macroperipatus to which P. ohausi was allocated. In 1975, Peck revised the New World Onychophora and considered Macroperipatus a valid genus of Peripatidae. In 1988, Read studied the integument of
Macroperipatus ohausi (Bouvier, 1900) and showed that the base of the primary papillae is rounded rather than quadrangular, as in Macroperipatus torquatus (von Kennel, 1883), type species of Macroperipatus. Furthermore, Read ]]>
In a recent study of Brazilian peripatid onychophorans, Oliveira, Wieloch & Mayer (2010) revised ]]>
Macroperipatus –M. acacioi and M. machadoi Oliveira & Wieloch, 2005– and also showed that the base of the dermal papillae of these species are rounded rather than quadrangular, according to their original descriptions (Marcus & Marcus, 1955; Oliveira & Wieloch,2005). In addition, the apical pieces of the primary papillae are well developed and, there- ]]>
Macroperipatus. Because of the presence of crural tubercles on only two pre-genital segments in males (Clark, 1913; Peck, 1975), the rounded dorsal papillae, and the low number of scale ranks in basal pieces of the primary papilla (Read, 1988), they included these two species in Epiperipatus.
As no specimen of M. ohausi had been collected in the last thirty years and its taxonomic status has not been reviewed, we redescribed and illustrated in this study its color in vivo, and the morphology of the dorsal integument using SEM. ]]>
Materials and Methods
Material: The two specimens were col- lected in the Reserva Particular do Patrimônio Natural dos ]]>
Fig. 1). In respect of syntypes, some characters of the syntype (female) housed in the onychophoran collection of the Zoolo-gisches Museum Hamburg (Hamburg, Germany) were verified by Hilke Ruhberg (personal communication). Repository acronyms are as follows: MNRJ-Museu Nacional/UFRJ, Rio de Janeiro, State of Rio de Janeiro, Brazil and ZMH-Zoologisches Institut und Zoologisches Museum, Hamburg, Germany. ]]>
Fixation: Specimen (MNRJ 0056) was fixed directly in alcohol 70%. Specimen (MNRJ0058) was previously anesthetized in ethyl acetate and then fixed in alcohol 70%, according to the protocol of Monge-Nájera & Morera-Brenes (1994). After fixation, the specimens were sexed and examined under a stereomicroscope and with scanning electron microscope (SEM). SEM methods included: (1) Cleaning: pieces were rehydrated and cleaned with a 3% solution of Sodium Hypochlorite (NaClO) and distilled water. They were then dehydrated using an increasing alcoholic series: 30%, 50%, 70%, 80%, 90% and 100%. (2) Critical point: all the examined pieces with SEM were critical point dried in model Baltec CPD 030, and then covered by a coat of gold (approximately 200Å=2.10-10m) in Denton Desk Iv gold sputter model.
Nomenclature: Dental formula and descriptive terminology follows Morera- Brenes & Monge-Nájera (2010) and Oliveira et al. (2010), respectively.
Color study: Pictures were taken with a Fuji Finepix S1500 (4-mega pixels) camera. Coloration was described based on living specimens. Following Kury & Orrico (2006), standard names of the 267 Color Centroids of NBS/ ISCC Color System (Mundie, 1995) were used in descriptions and given between parentheses.
Results
Family Peripatidae
Genus Epiperipatus Clark, 1913
Epiperipatusohausi (Bouvier, 1900) new combination
Macroperipatus ohausi: Peck 1975: 347; Sampaio-Costa et al. 2009: 558; Oliveira et al. 2012:16.
Fig. 2-12.
Type material: (Syntypes: two females (ZMH and MNHN), BRAZIL, Petrópolis, 31. I. 1900, Dr. Ohaus leg. (Not examined).
Additional material examined: BRAZIL, state of Rio de Janeiro, Nova Iguaçu, Reserva Particular do Patrimônio Natural dos Petroleiros, 23.XII.2009, Guipponi, A.L.; Sampaio-Costa, C. leg. (1 female, MNRJ 0056); idem, 11.III.2010, Chagas-Jr, A.; Giupponi, A.L.; Kury, A.; Sampaio-Costa, C. leg. (1 male, MNRJ 0058).
Diagnosis: Antennae with 38 to 41 complete rings; outer jaw blade composed of one long curved principal tooth and an additional accessory tooth; inner jaw blade with one principal tooth, one accessory tooth, a diastema, and a series of 12 denticles; dorsal papillae rounded; basal piece of dorsal primary papillae conical; basal and apical pieces of dorsal pri- mary papillae with 7 to 8 and 3 to 4 scale ranks, respectively; leg pairs 27 to 29; ventral organs visible; anal glands of male indistinguishable.
Redescription
Measurements: Female ]]>
Color (for living specimens): Body background brownish gray (64) with dorsal light spots (primary papillae) pinkish gray (10). Dorsomedian furrow is pale violet (214), with light olive gray parallel diamonds (112). The head is moderate brown (58) and antennae are grayish to reddish purple (245) (
Fig. 2). Dorsal side of legs is grayish to purplish pink (253), ventral side is pale purple (227), and the soles are yellowish white (92). The ventral side is dark pink (6) with light pink (4) ventral organs and light spots pinkish white (9), randomly dis- tributed (Fig. 3). One syntype (ZMH) is totally discolored.
Integument: Twelve plicae per segment, seven of these cross over to ventral side. The incomplete plicae are interrupted over the legs (Fig. 6).
Hyaline organs: On dorsal side, between plicae and parallel to the dorsomedian furrow (Fig. 7).
Dorsomedian furrow: Slender, and starts at mid-point of head and finishes near anus.
Dorsal papillae: Female (MNRJ 0056). The dorsal papillae are close to each other and the size varies. ]]>
Fig. 6). The primary papillae are the biggest dorsal papillae (Fig. 6, 8 and 9); they are rarely in sequence or separated by accessory papillae. The number of accessory papillae varies between two primary papillae. The longest primary papillae are discolored with base diameter larger than plicae. These papillae are always found on alternate plicae in “longitudinal lines” present on dif- ferent aspects of dorsal side. Depending on its relative position to the dorsomedian furrow, these ]]>
The conical basal piece of the primary papillae has seven (Fig. 8) or nine scale ranks. But nine scale ranks were observed only in the female (MNRJ 0056). These scales are lanceolate and increase in length near the top (Fig. 8). Constriction ]]>
Fig. 8 and 9). The apical piece has three scale ranks on posterior side and four on anterior side.
The accessory papillae are conical ]]>
Fig. 6). The scales on the bottom are more elongated than on top.
Head. Without evident structures. Antennae. 38 to 41 complete rings. One syntype (ZMH) has 38 rings. In ]]>
Frontal organs. Present at antennal base on ventral side with length about equal to four fused antennal papillae. Eyes. One on each antennal base. Mouth parts. Opening surrounded by one anterior oral lobe (unpaired lip) and seven pairs of oral lips. In each pair of oral lip there is a median series of small spines. One syntype (ZMH) has ]]>
Jaws. (MNRJ 0056). Outer jaw blade. Composed of one long curved principal tooth and an additional accessory tooth (formula: 1/1) (Fig. 4). The accessory tooth is knife-shaped, curved at the top edge and straight at the bottom edge. Inner jaw blade. With one ]]>
Fig. 4 and 5). The accessory tooth of both blades is similar in shape.
Legs: Female with 29 and male with 28 pairs of legs. Spinous ]]>
Fourth and fifth legs have four spinous pads with similar width, and length increases between the first three pads (Fig. 10 and 11). The length of the fourth spinous pad is similar to that of the second. Nephridial tubercle. Present on fourth and fifth pairs of legs, between 3rd and 4th spinous pads. It is totally connected with the third pad, articulating with the fourth, but is free from it (Fig. 10 and 11). ]]>
Foot papillae. Feet have two anterior and one posterior foot papillae. ventral foot surface with six bristles: two pairs in proximal setiform ridge, and at least one bristle on each distal setiform ridge (Fig. 12). Ventral organs. They are clearly visible in vivo. Post-fixation they are represented by depressions on the ventral integument.
Genital opening. Located between penultimate pair of legs in both sexes, but the morphology of the genital opening is different among the sexes. The genital opening in female is poorly preserved, but it seems to have two crossed furrows; the transversal one is only weakly visible. Moreover, the male has a single long furrow parallel to the main body.
Male characters: Crural papillae are pres- ent in one pre-genital pair of legs (a single tubercle per leg). Anal glands are only recog- nized by their corresponding open pores.
Type locality: Petrópolis, state of Rio de Janeiro, Brazil.
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Distribution: State of Rio de Janeiro (Petrópolis and Nova Iguaçu, new record), Brazil.
Discussion
The Neotropical ]]>
Peripatus,
Epiperipatus, Macroperipatus, Speleoperipatus Peck, 1975 and Plicatoperipatus Clark, 1913, but the presence of eyes and 12 plicae per segment distin- guishes them from Speleoperipatus and
Plicatoperipatus, respectively.
Because of the conical basal piece with a roundish base and the spherical, asymmetrical and robust apical piece, the primary papillae of the analyzed specimens suggest a higher resemblance to Peripatus and Epiperipatus than to Macroperipatus. According to Read (1988), the quadrangular shape of dorsal papillae seen in M. torquatus is caused by the outline grooves. In addition, the scales of the basal piece are not arranged in ranks in this species (Oliveira et al., 2010). Therefore analyzed specimens do not belong to Macroperipatus.
The dorsal integument of all Macroperipatus species is not known in detail, although according to Read (1988) the dorsal primary papillae of M. acacioi, M. ohausi and M. guianensis (Evans, 1903) are rounded and contrast with quadrangular primary papillae of M. torquatus. In a recent review of two Brazilian ]]>
Macroperipatus–M. acacioi and M. machadoi– Oliveira et al. (2010) also showed that in these species the bases of the dermal papillae are roundish and covered with ]]>
Epiperipatus. They also suggested that the dorsal papillae should be reinvestigated in other species of Macroperipatus by SEM. ]]>
Read (1988) registered one M. ohausi specimen, collected in 1981 by J. C. Diniz and C. R. Brandão, in Petrópolis. We contacted C. R. Brandão and he informed us of the exact collection point where the specimen was sampled (C. R. Brandão pers. comm.). We returned several ]]>
The two new specimens were collected 28km from Petrópolis, type locality of M. ohausi
(Fig 1). The characters present in these specimens agree with the description of M. ohausi sensu Bouvier (1900, 1905), except for the number of leg pairs. The number of leg pairs is not fixed in Neotropical Peripatidae species (Bouvier, 1905; Campiglia ]]>
M. ohausi, based mainly on the description of the primary papillae morphology. Then, Read (1988) included the number of scale ranks in the description of this species based on recent specimens collected in 1981 by J. C. Diniz and C. R. Brandão in Petrópolis.
The dorsal primary papillae described by Read (1988, figs. A-D) were compared to the specimens from RPPN dos Petroleiros and also agree with them in morphology. The morphology of the dorsal primary papillae and the range of leg pairs show that Epiperipatusohausi is more closely related to the species of Epiperipatus from the state of Minas Gerais (Southeastern Brazil): E. acacioi, E. machadoi, ]]>
E. diadenoproctus Oliveira, Lacorte, Fonseca, Wieloch & Mayer, 2011; E. adenocryptus Oliveira et al. 2011; and E. paurognostus Oliveira et al. 2011. Epiperipatusohausi
has seven or eight scale ranks on the base of the primary papillae, while E. acacioi, E. machadoi, E. diadenoproctus, E. adenocryptus have 5-6, 5-8, 5-8 and 5-7, respectively. Four scale ranks on E. adenocryptus, three on E. ohausi
and E. acacioi, and two on E. paurognostus were counted from the posterior side of the apical piece, but for E. diadenoproctus it was not described.
Epiperipatusohausi, E. ]]>
and E. machadoi have the asymmetrical apical piece in common, though it is spherical in the first species and ovate in latter two (although some E. acacioi specimens have a ]]>
E. acacioi, E. machadoi and E. ohausi. However, it is spine-shaped in the cryptic species from Minas Gerais (Oliveira et al., 2011). The range of leg pairs is different among these six Brazilian Southeastern species. In increasing order, the ranges are: 27 to 29 in E. ohausi; 25 to 28 in E. ]]>
(n=34); 26 to 29 in E. paurognostus (n=15, see Oliveira et al., 2011); 26 to 30 in E. adenocryptus (n=25, see Oliveira et al., 2011); E. diadenoproctus (n=62, see Oliveira et al., 2011), and 27 to 31 in E. machadoi (n=21). The anal glands are absent in males of E. ohausi
, but the respective pores are present. These pores are observed on the ventral board of the anus. In E. acacioi and E. machadoi the anal glands seem also to be absent, while in E. adenocryptus, E. diadenoproctus and E. paurognostus these structures are present in the ventral board of the anus. The SEM primary papillae ]]>
Fig. 5, 7 and 8) analysis of M. ohausi showed that the base is not square-shaped as in M. torquatus, but ]]>
Epiperipatus.
The new record of E. ohausi has ]]>
Acknowledgments
We are ]]>
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1. Departamento de Biologia e Zoologia, Instituto de Biociências, Universidade Federal de Mato Grosso. Avenidan Fernando Correa da Costa, 2367, Boa Esperança, CEP-78060-900, Cuiabá, MT, Brasil; amazonaschagas@gmail.com 2. Departamento de Zoologia, Instituto de Biociências, Universidade de São Paulo. Rua do Matão-Travessa 14, nº 321, Cidade Universitária, São Paulo, SP, Brasil. CEP 05508-090; csampaioc@gmail.com
Received 20-vIII-2013. Corrected 06-Iv-2014. Accepted 05-v-2014.
